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Rivista Italiana di Paleontologia e Stratigrafia volume 114 no. 1 1 pl. pp.

119-132 March 2008





Received: May 10, 2007; accepted: October 16, 2007

Key words: Stratigraphy, Vertebrate fauna, Testudo, Pliocene, letteratura alla ``fauna di Mandriola'', che viene qui rinominata ``Fauna
Mandriola, Capo Mannu, Sardinia. locale Capo Mannu D1''.
I vertebrati della fauna Capo Mannu D4 appartengono a rettili
Abstract. In the North-West Sinis Peninsula (Western Sardinia) (Ordine Chelonii) e mammiferi (Ordine Artiodactyla, famiglie Bovidae
a stratigraphic sequence, the Capo Mannu Fm., that evolves from e Suidae). Il fossile di testuggine mostra caratteri che consentono la
marine-littoral to continental-dunar, is present. A vertebrate fauna is definizione di una nuova specie. Il suide eÁ rappresentato da reperti
reported within the middle part of this coastal dune complex. These che indicano la sua appartenenza alla specie endemica Sus sondaari. I
new remains, here referred to as the Capo Mannu D4 Local Fauna, bovidi sono il gruppo meglio rappresentato, benche i resti siano piut-
are slightly younger in the sedimentary succession than the Local tosto frammentari. Essi appartengono almeno a due forme distinte. Una
Fauna known in the literature as Mandriola and here re-named the di queste eÁ confrontabile al genere Nesogoral, che rappresenta uno degli
Capo Mannu D1 Local Fauna. elementi piuÁ caratteristici della fauna endemica sarda attribuita al co-
Vertebrates from Capo Mannu D4 include reptiles (Chelonii) siddetto complesso a ``Nesogoral'', che documenta un intervallo di tem-
and mammals (Bovidae and Suidae). The tortoise fossil shows peculia- po dal Pliocene superiore alla parte iniziale del Pleistocene inferiore.
rities of shape that require the naming of a new species. The suid L'associazione di Capo Mannu D4 rappresenta quindi una delle piuÁ
material includes a fragmentary palate and a partial mandible, referable antiche segnalazioni della presenza del complesso a Nesogoral nell'am-
to the endemic species Sus sondaari. Bovids are well represented in the biente insulare Sardo.
assemblage, although quite fragmentary. They have been identified as
belonging to at least two forms. One of these is comparable in mor-
phology to Nesogoral, one of the most characteristic taxa of the Late Introduction
Pliocene-Early Pleistocene Sardinian endemic fauna (the so called ``Ne-
sogoral complex''). On the whole, the Capo Mannu D4 assemblages A small vertebrate fauna was found within an
open a new window on the Plio-Pleistocene terrestrial faunas of the interdune deposit at the base of the Capo Mannu For-
Sardinian Island.
mation, overlying the Mandriola Limestone (North-
Riassunto. Nella penisola del Sinis (Sardegna Occidentale) affio- West Sinis, Fig. 1). These remains were first reported
ra la formazione Pliocenica di Capo Mannu, che eÁ caratterizzata da by Pecorini et al. (1974) as Middle Pliocene in age.
un'alternanza di corpi dunari e paleosuoli ed evidenzia un'evoluzione Other vertebrate remains found in the dune com-
paleoambientale da marino-litorale a continentale. Nella parte interme- plex of the Capo Mannu Formation have been recov-
dia di questo complesso dunare sono stati rinvenuti resti fossili di verte-
brati di ambiente terrestre. Questi resti vengono attribuiti, nel presente
ered by two of us (SC and LL). These new remains were
lavoro, alla ``Fauna locale Capo Mannu D4'', in riferimento all'unitaÁ included in a new stratigraphic reconstruction of the
dunare in cui sono stati rinvenuti. Da un livello, stratigraficamente al Mandriola-Capo Mannu succession in a previous paper
di sotto della fauna Capo Mannu D4, provengono i fossili riferiti in (Carboni & Lecca 1995).

1 * Corresponding author. Dipartimento di Scienze della Terra, UniversitaÁ degli Studi di Firenze - Via G. La Pira 4, 50121 Firenze.
2 Dipartimento di Scienze della Terra, UniversitaÁ degli Studi di Cagliari - Via Trentino 51, 09127 Cagliari.
120 Abbazzi A., Carboni S., Delfino M., Gallai G., Lecca L. & Rook L.

Upper Pleistocene beach sandstones and conglomerates

referred to the Tyrrhenian transgression (Marine Isoto-
pic Stage, MIS 5e) and continental talus referred to the
Marine Isotopic Stages MIS 4-2 (Carboni & Lecca 1985,
1995; Lecca & Carboni 2007).
Mandriola Limestone
The Mandriola Limestone outcrops at the edge of
the North-West Sinis area. It has been referred to the
Lower Messinian as ``Calcari Laminati del Sinis'' by
Cherchi et al. (1978). Seismostratigraphic studies on
Pliocene hemipelagic marl sampled on the continental
shelf (Lecca et al. 1986), suggest that the Mandriola
Limestone is stratigraphically situated a few tens of
metres above the Pliocene hemipelagic marl. Biostrati-
Fig. 1 - Location of Capo Mannu and Mandriola sections. A-B: graphic studies recognized that this marl belongs to the
location of the cross-section shown in Fig. 3. Discoaster tamalis Zone, Reticulofenestra pseudoumbili-
ca Subzone, NN 15 (approximately 3.5 Ma; Francolini
The aim of this paper is to describe in detail the et al. 1990). The Mandriola Limestone is mainly consti-
unpublished vertebrate remains of the Capo Mannu tuted by a foraminiferal grainstone with large ripple
Fm., as well as to provide a detailed stratigraphic con- cross-laminations and hummocky structures of shore-
text for this dune complex and the vertebrates occurring face environment. This unit, about 10 m thick, gently
therein. dips westward and outcrops for approximately 1.5 km.
The upper part contains several storm-related levels that
are very rich in pelecypods. Within these levels are in-
terbedded terrigenous sediments containing sparse
rounded clasts of Messinian limestones (``Calcari Lami-
The Pliocene-Pleistocene sequence in the north- nati del Sinis'') and alkaline basalts. The origin of the
ern Sinis (Mandriola Limestone and Capo Mannu Fm.) volcanic clasts may be individuated in the Montiferru
overlies (with a possible erosional unconformity) a and Sinis areas, where the radiometric ages of related
Middle Miocene-Messinian sequence resting on the Oli- volcanites are 3.9 Ma (``lower basanites'' of the Monti-
go-Miocene volcanic basement (Fig. 2). The Pliocene- ferru volcanic complex), 3.12 Ma (basalts of San Salva-
Pleistocene sequence can be summarized as a shoreface tore di Sinis) and 2.8-3.0 Ma (basalts of the southern
unit followed by various dune units, locally overlain by side of the Montiferru volcanic complex; Beccaluva et

Fig. 2 - Synthetic cross-section of the Sinis (Capo San Marco - Capo Mannu) stratigraphic units and location of faunal deposits.
Fossil vertebrates from Capo Mannu, with description of a new Testudo species 121

Fig. 3 - Capo Mannu dune complex upper part cross-section and detailed location of collected fossil remains.

al. 1985). On the basis of the full data set the above- ing 10 cm thick beds, mainly consisting of up to 0.5 cm
mentioned sedimentary shoreface sequence is to be in- fragments of pelecypods alternated with level of similar
terpreted at the Zanclean-Piacenzian transition (Carbo- thickness consisting of finer bioclastic sediments. Such
ni & Lecca 1995). units display a significant lateral continuity and an im-
portant aggradational component, and thus form tabu-
The dune complex of the Capo Mannu Formation lar accretions which are on average 10-12 m thick.
The Capo Mannu Formation (CMF), primarily In the dune subunit characterized by the finer
defined by Pecorini et al. (1974), is a calcareous and bioclastic facies, the main depositional structures are
terrigenous 50 m thick sand body, made up of four represented by discontinuous compound sets of cyclic
staked and laterally continuous dune units, overlain crossbeds and reactivation surfaces of low angle. The
by three other dune units of lesser lateral continuity. accretion is highlighted by prograding parallel laminar
On the whole, at least nineteen dune subunits can be sets tending to sigmoidal shapes. These structures devel-
identified in the aeolian complex (Carboni & Lecca op laterally by moderate fluctuations in the flow direc-
1995; Fig. 2). tion, ranging from south-eastward to north-eastward,
The hierarchy of the depositional discontinuities producing some herringbone structures.
shows internal boundaries of 3rd, 2nd, and 1st order sensu The bioclast-rich pelecypod subunit displays an
Brookfield (1977). The 1st order boundaries, generally oblique parallel tabular set 3-4 m thick developed
represented by reddish clayey-silty sediments, may be roughly south-eastward at angles of up to 30ë. It must
associated both with interdune deposits and with inter- be noted that in the interdune levels and within the
ruptions of the eolian processes caused by climate massive strata of the D3 and D4 dune units, an exemplar
change. of the tortoise (Tegas 2000) and some of the vertebrate
The first dune unit (D1), at the base of the CMF, remains (described below) have been found (Figs. 2, 3).
consists of slightly dipping foraminiferal aeolian banks The last three dune units (D5, D6 and D7), in the
with at least two interbedded dump or clayey pond upper part of the dune complex, are laterally discontin-
deposits at the base of the lee side of the dune. The Capo uous and also composed of calcareous sandstones with
Mannu D1 local fauna (also known in the literature as foraminifers with variable terrigenous contents. Their
Mandriola; see below) derives from the first clayey de- internal geometry is more varied and usually shows
posit (Fig. 2). This unit extends throughout the coastline tabular and low-angle sigmoidal prograding coset, low
between Mandriola and the Capo Mannu promontory, planar parallel in the higher parts. In these units the
with a total thickness of approximately 3-4 m. components are lithic-quartzose and bio-detrital
The next three overlying dune units (D2, D3 and (benthic microfauna).
D4) are developed along the entire Capo Mannu cliff. The 1st order boundaries with reddish clayey-silty
They exhibit distinct structuring which often differs sediments separate the main seven units, and minor un-
markedly in grain size, composition, textural features conformities indicate several dune subunits within each
and degree of diagenesis, and they display numerous main unit. These reddish sediments show a quartzose
discontinuities of lower order in the accretion architec- terrigenous component with homogenous fine sand-silt
ture, in places with abundant root traces. The composi- grain-size. The clay percentage varies from entirely
tion of such units varies both laterally and vertically, clayey laminae to calcareous and iron/manganese films.
shifting from levels with high content of brackish The reddish sediment layers are up to 2 m thick, but
benthic foraminifers (Cibicides, Elphidium, Rotalinidae, they tend to thin laterally. Generally, the higher part
Miliolidae) and Ostracoda to levels with a dominant contains abundant root traces, which may contain ver-
terrigenous siliciclastic component. In some units, a tebrate bones and shells of Gastropoda Pulmonata. The
bioclastic component of pelecypods is dominant, form- very sandy sediments of the first-order unconformities
122 Abbazzi A., Carboni S., Delfino M., Gallai G., Lecca L. & Rook L.

Fig. 4 - Chronological scheme and

correlation of Capo Mannu
local faunas with the bio-
chronological mammal u-
Fossil vertebrates from Capo Mannu, with description of a new Testudo species 123

must be attributed to re-sedimentation of finer grains Some of the early remains were sent by G. Pecor-
derived from large dunes of a low energy aeolian regime ini to P.L. Ambrosetti (Perugia) for taxonomic and bio-
in a wet climate. chronologic study, but they were never studied in detail.
Thanks to the kind help of S. Gentili (Perugia), these
remains (including a fragmentary skull and mandible of
Fossil vertebrates from the Capo Mannu Formation a small suid from D4, and a mandible and a few isolated
teeth of a bovid from D1) have been re-located and
Capo Mannu D1 local fauna: a debated chronological made available for the present study, along with the
attribution entire collection of the Capo Mannu D4 fauna.
The occurrence of a small vertebrate assemblage in The fossils here described are housed in the Earth
the interdune damp or pond deposits at the base of the Sciences Department of Cagliari University. The acro-
Capo Mannu Fm. (Fig. 2, ``Mandriola deposit'') was ori- nym ``GP'' refers to the early collection by G. Pecorini,
ginally reported by Pecorini et al. (1974). These authors while the acronym ``CL'' refers to the recent collections.
attributed the vertebrate assemblage to the Middle Plio-
cene on the basis of biochronologic interpretations.
This small assemblage has been reported in the Systematic Paleontology

literature with different names. While most authors

use the hamlet name ``Mandriola'', others (e.g. van der Reptilia
Made 1988, 1999) refer to it as ``Capo Mannu''. In order
to avoid confusion with the name of the Lower to Mid-
dle Pliocene Mandriola Limestone, we propose here to Chelonii Brongniart, 1800
name all the vertebrate fauna from the dune complex of Testudinidae Batsch, 1788
the Capo Mannu Formation as Capo Mannu. Thus, the Testudo Linnaeus, 1758
Mandriola mammal assemblage of previous authors will
be referred to as the Capo Mannu D1 Local Fauna.
Some authors (Esu & Kotsakis 1979, 1983; Kot- Testudo pecorinii Delfino n. sp.
sakis 1980; Esu 1984), mainly on the basis of regional Holotype: CL 102, a nearly complete shell housed in the Earth
correlations, assigned the Capo Mannu D1 local fauna Science Department of Cagliari University, Italy (Figs. 5,6).
to MN 14 mammal unit of the early Ruscinian (Early Etymology: the species is dedicated to Prof. G. Pecorini, hon-
Pliocene; Fig. 4), while others (e.g. van der Made 1988) ouring his outstanding contribution to the study of Sardinian geology.
Type locality: Capo Mannu (San Vero Milis Municipality); base
agreed with the original (Pecorini et al. 1974) attribution of D4 dune of the Capo Mannu Formation; Upper Pliocene.
to the Middle Pliocene. Diagnosis: the only fossil or extant member of the genus Tes-
The above mentioned biostratigraphic characteri- tudo, group Testudo sensu stricto (therefore with a hinge between hy-
zation of the Pliocene Mandriola Limestone (Francolini poplastra and xiphiplastra), characterized by five evident bosses sagit-
tally aligned on the carapace; the bosses, developed in correspondence
et al. 1990), constrains the age of the Capo Mannu D1 to the central region of the five vertebral scutes, are medio-laterally
local fauna as corresponding to the time interval identi- elongated, apically convex and well separated from each other.
fied by marine biostratigraphy as the NN15/NN16
Zones (Fig. 4), in contrast with its attribution to the Description
early Ruscinian (MN14; Kotsakis 1980; Esu & Kotsakis General preservation, shape and size. The shell is
1983; Esu 1984). filled with matrix and is rather complete in terms of
Recently, Angelone & Kotsakis (2001), on the preserved skeletal elements, but even if it retains the
basis of a new study of the Capo Mannu D1 murid overall morphology, the bone surface displays only
(Raghapodemus azzarolii), assigned this fauna to the few anatomical details. The original surface morphol-
late Ruscinian. Therefore, they also correlate the age ogy, with furrows corresponding to the limits between
of this deposit with the Zanclean-Piacenzian transition, the horny scutes, is preserved only in the antero-median
confirming the reconstruction of Carboni & Lecca region of the carapace and in a small antero-lateral cor-
(1995). ner of the plastron. A large portion of the bones (whit-
ish in colour) has lost the outer layers and shows the
The vertebrate remains of the Capo Mannu D4 Local eroded spongy inner tissue or is covered by a thin
Fauna brownish arenaceous matrix.
Geological/sedimentological surveys of the Capo The best-preserved area of the carapace is the
Mannu dune complex have recovered a number of ver- medial part, since the anterior region is eroded to some
tebrate remains, in early times by Prof. G. Pecorini, and extent and the posterior area is heavily damaged and
later by two of us (SC and LL), from a fossiliferous level incomplete (the posterior peripherals are only partly
corresponding to the base of the D4 dune (Fig. 3). preserved). The plastron is nearly complete. A small
124 Abbazzi A., Carboni S., Delfino M., Gallai G., Lecca L. & Rook L.

Fig. 5 - Testudo pecorinii Delfino n. sp. - Photographs and interpretative drawings of the shell in left lateral, upper and lower view. Dark grey
indicates the matrix filling the shell, light grey indicates the thin concretion. The sutures involving neural elements have not been
represented in lateral view. Scale: 10 cm.
Fossil vertebrates from Capo Mannu, with description of a new Testudo species 125

portion of the surface of the fourth neural and the not occupy the entire vertebral surface but only its cen-
fourth right pleural is collapsed inward. tral area (therefore protuberances are well separated
A possible bony structure of unknown origin (not from each other) and are medio-laterally elongated.
necessarily belonging to the tortoise) is sealed by the On both sides, the first pleurals preserve only the pos-
matrix on the carapace in the region of the posterior- terior suture; the sutures delimiting the following two
most right pleural. pleurals on the right and three on the left side are rela-
As for the general shell shape, it is remarkable tively visible: these trapezoidal elements show a typical
that, in dorsal view, the central region of the carapace alternation, with the widest base alternately placed dor-
(whose peripherals are engaged in the bridge) is rather sally and ventrally. Only the dorsal end of the pleurals
rounded and separated from the anteriormost area participates in the protuberances generated by the neur-
(whose peripherals are free) by a slight constriction that als. The peripherals are better preserved on the left side,
is particularly evident on the left side; the development where at least five elements show clearly perceivable
of the anterior area, and therefore the origin of such a sutures, and they are rather large antero-posteriorly.
constriction, is not known, since the peripherals are The horny mosaic is represented by the scute sulci
distally eroded or broken (mainly on the right side). of the second and third vertebrals (whose boundaries are
A hint of a posterior constriction, corresponding to not completely visible) and by the costals and marginals
the boundary between the peripherals engaged in the corresponding to the preserved bony plates. In dorsal
bridge and those constituting the rim of the posterior view, the vertebral scutes are wider than the costal ones.
shell opening, seems to be present, but the development In anterior and, to a lesser extent, posterior view, the area
of the posterior area of the shell cannot be evaluated delimited by the vertebrals corresponds to a convexity of
because of the poor preservation. the shell outline delimited by a depression along the fur-
Even if a weak dorso-ventral deformation cannot rows between vertebrals and costals. In the proximal
be ruled out, the anterior opening of the shell is rather sector of each costal a hint of a boss is perceivable in some
narrow. The moderate concavity of the plastron could cases. Sutures between pleurals and peripherals corre-
be related to such a possible weak deformation or re- spond to the furrows between costals and marginals.
present a male character. Plastron. The morphology of the plastron is much
The shell has a total length of 22.5 cm, a maximum less interpretable than that of the carapace. Except for a
width of 17.8 cm and a maximum height of 12.2 cm. short tract of the furrow between right gular and hum-
Carapace. The first neural is barely visible but eral, no furrows left by the horny shields are visible on
seems to be distinctly elongated and approximately rec- the plastron. The epiplastra show a thick dorsal lip
tangular; it is followed by an octagonal element and whose morphology cannot be evaluated in detail be-
then by a rectangular one (whose antero-posterior cause of the presence of matrix filling the shell; the
length is greater than its width). The neurals are mark- epiplastral area covered by the gular horny shields pro-
edly convex and concave: such a morphology confers to trudes markedly anteriorly and represents a concavity
the shell dorsal outline a ``peaks and valleys'' pattern, of the ventral plastral surface. The entoplastron seems
with peaks approximately corresponding to sutures be- to be rather rounded. The gular shields penetrated into
tween adjacent bony neurals (and the centre of the the entoplastron (but due to the presence of matrix it is
horny vertebral shields) and valleys to the neural surface not possible to assess how deeply they penetrated). The
(and therefore the furrows representing boundaries be- xiphiplastra are still joined together but they are dis-
tween adjacent shields); the protuberances have a uni- tinctly separated from the hypoplastra by a gap of some
formly convex apex (they are not apically flattened), do millimetres: they have not been significantly dislocated

Fig. 6 - Testudo pecorinii Delfino n. sp. - Shell in anterior and posterior view. Scale: 10 cm.
126 Abbazzi A., Carboni S., Delfino M., Gallai G., Lecca L. & Rook L.

thanks to the presence of the matrix. The boundaries and possibly in the Early Pliocene of Tunisia (see refer-
between elements of the carapace and plastron not men- ences in Lapparent de Broin 2000; 2001; 2002).
tioned here are not visible. The Sardinian land tortoise fossil record is limited
Discussion. Due to the poor preservation, the to a partial shell from the Plio-Pleistocene locality of
morphology of all the bony plates and horny shields Orosei (Abbazzi et al. 2004) referred to T. cf. T. her-
of the shell cannot be assessed. However, the morphol- manni, and to a few Holocene Testudo remains from
ogy and relationships of the available elements, chiefly archaeological contexts (cfr. Delfino 2002).
the shape alternation in the neurals and pleurals, as well At present, three tortoises inhabit Sardinia, T. her-
as the correspondence of sutures between pleurals and manni, T. graeca and T. marginata, but only the first one
peripherals to the furrows between costals and margin- has been considered autochthonous (Amori et al. 1993;
als, and the general architecture of the shell, are consis- Lanza & Corti 1993). However, the presence of a mor-
tent with that of a member of the genus Testudo. The phologically peculiar Testudo s.s. in the Late Pliocene of
separation of the xiphiplastra from the hypoplastra in- Capo Mannu does not allow us to extrapolate a genetic
dicates the presence of a hinge allowing movements of continuity with the extant Sardinian populations be-
the posterior lobe of the plastron: such a characteristic longing to this group, T. graeca and T. marginata.
identifies the Testudo sensu stricto group, represented by Concerning the origin of the tortoises of Sardinia,
T. graeca and other extant forms inhabiting the Medi- the analysis of mitochondrial RNA recently suggested
terranean region (see Lapparent de Broin 2000 for a that living populations of T. hermanni (as well as all the
list). The fact that the movable xiphiplastra were not Testudo spp.) on Sardinia and other Mediterranean Is-
dispersed prior to fossilization, suggests rapid burial lands have been introduced by man (Van der Kuyl et al.
2002). Such a conclusion is not supported by the fossil
of the shell before soft tissue decomposition. A further record which shows the presence of T. hermanni in the
character shared with T. graeca (Amiranashvili 2000), Plio-Pleistocene of Sardinia (Abbazzi et al. 2004) and,
and other species of its group, is the width of the ver- above all, in the Middle Pleistocene of Corsica (Hervet
tebrals when compared to the costals. & Salotti 2000; Hervet 2001). If a gap in the continuity
The specimen from Capo Mannu is characterized between these Pleistocene occurrences and the extant
by a peculiar morphology not known in any fossil or populations is present, it is necessary to hypothesize
recent member of genus Testudo s.s.: the carapace sur- that the introduction operated by man followed (or
face is distinctly raised in correspondence with each of caused) the extinction of the autochthonous populations
the vertebral scutes. Such a morphology is quite differ- on both islands.
ent from that sometimes occurring in wild Testudo po-
pulations (see Guyot Jackson 2004) or originating in Mammalia
captivity from an unbalanced diet, such as the calcium
/ phosphorous unbalanced ratio that causes a morpho- Artiodactyla Owen, 1848
logical anomaly known as ``pyramiding'' (in which, Suidae Gray, 1821
since the entire vertebral surface is convex, the bosses
are contiguous and, moreover, usually apically flat- Suinae Gray, 1821
tened), but it is somehow reminiscent of the condition Sus Linnaeus, 1758
seen in some species of the genus Psammobates or
``Geochelone''. Sus sondaari van der Made, 1999
A weak development of dorsal protuberances is
seen also in Testudo amiatae Pantanelli, 1892, from 1980 Sus scrofa ssp. Ambrosetti et al., pag. 247.
the Upper Miocene of southern Tuscany, but this spe-
cies is characterized by xiphiplastra that are firmly su-
tured to hypoplastra and therefore cannot be ascribed to
the Testudo s.s. group. PLATE 1
On the basis of this peculiar set of characters we Sus sondaari - A, B occlusal and labial views of GP2 incomplete
propose a new species, Testudo pecorinii, belonging to maxillary from the Capo Mannu D4 local fauna, and C, D, E occlusal,
lingual and labial views of GP1 fragment of left hemimandible with
the Testudo s.s. group. M3-M1 from the Capo Mannu D4 local fauna.
The presence of a Testudo s.s. in the Late Pliocene Nesogoral sp. - F, G, H labial, lingual and occlusal views of mandible
of Capo Mannu represents the first fossil evidence for GP1 from the Capo Mannu D1 local fauna, and I, L, M labial, lingual
this group in peninsular and insular Italy. This group and occlusal views of CL127 fragment of left hemimandible with M2
and erupting M3 from the Capo Mannu D4 local fauna. Small sized
probably evolved in Europe since it can be traced back bovid of uncertain attribution - N, O, P occlusal, labial and lingual
to the Turolian of Pikermi, Saloniki and Lesvos Island, views of CL 137 right hemimandible from Capo Mannu D4 local
but it is already present in the Late Pliocene of Morocco fauna.
Fossil vertebrates from Capo Mannu, with description of a new Testudo species 127
128 Abbazzi A., Carboni S., Delfino M., Gallai G., Lecca L. & Rook L.

1988 Sus aff. nanus van der Made - pag. 373, Plate 3, Figs 7-9. the fragmentary status of most specimens does not al-
1999 Sus aff. sondaari van der Made, pag. 347. low for a firm taxonomic attribution.
Material: Capo Mannu D4 local fauna - GP3, palate; GP4, The greater part of the dental remains are referred
fragment of mandible; CL123c, right I3; CL106, molar fragment. to the small sized bovid Nesogoral sp., this genus being
one of the most typical elements of the Sardinian en-
Remarks. The palate is poorly preserved and demic fauna of Pliocene-Early Pleistocene age. A few
shows only the left M3, a small fragment of the left remains attest to the occurrence of another taxon, sig-
M2, and the right M2-M3 (Plate I, figs. A, B). The M2 nificantly larger than Nesogoral sp. (Bovidae gen. et sp.
crown is totally worn. The M3 is also affected by wear, indet., see below). Moreover, it is worth underlining
but it is still possible to observe the position of the main that among the fossils of similar size to Nesogoral sp.,
cusps and the enamel thickness. some dentitions do not fully conform to the description
Only the posterior part of the left mandible is of the latter taxon (cf. Gliozzi & Malatesta 1982); there-
preserved (Plate I, figs. C-E). It shows an incompletely fore, they will be considered separately.
erupted M3 and an M2. The angle between the horizon- The present study confirms a complex scenario,
tal and ascending rami of the mandible seems to be regarding the dispersal and evolution of Sardinian bo-
smaller than in recent wild pigs (Sus scrofa) but this vids in the insular environment, which has been already
could be due to diagenetic deformation. put in evidence by recent investigations. Indeed, the
No inferences can be added regarding I3 (crown occurrence of a diverse bovid fauna, with more than a
completely worn) and the small molar fragment. single species of Nesogoral and with another bovid
The dentitions of these new specimens from Capo genus has been reported in the Early Pleistocene fissure
Mannu D4, although smaller in size (Tab. 1), are similar infillings at Monte Tuttavista, Orosei (Abbazzi et al.
to the holotype of Sus sondaari from Capo Figari (van 2004; Palombo et al. 2006). Van der Made (2005) defined
der Made 1988; 1999) in basic morphology, crown the new species Nesogoral cenisae from the Middle
height and enamel thickness. Pleistocene of Campidano.
In the past, three other specimens (a right I2, a left
I , and a right P2) from Capo Mannu have been col-
lected in the Capo Mannu D1 interdune by a team of Nesogoral sp.
the Instituut voor aardwetenschappen of Utrecht (The
Netherlands; IVAU) during the operation of washing Material: Capo Mannu D1 local fauna - GP1, fragment of left
sediments in search of micromammals (van der Made, hemimandible with M3-M1; GP2, right M1-2.
Capo Mannu D4 local fauna - CL127, fragment of left hemi-
pers. com.). These isolated teeth (kept in the IVAU col- mandible with M2 and erupting M3; CL138, fragment of right mandible
lections) have been studied by van der Made (1988) and with M1-D2; CL123a, left M2; CL120 left M3.
attributed to Sus aff. sondaari. Van der Made, observing
in particular the wear of P2, noted that the Capo Mannu Remarks. The occurrence of Nesogoral in the de-
suid was a more primitive form because of the unre- posits of Capo Mannu (D1 local fauna) was reported by
duced premolar row. van der Made (1999), who described a few fossils in the
In fact, given the small size of the new Capo IVAU collections, Utrecht. The dentition here consid-
Mannu D4 remains, it is likely that the latter had al- ered is characterised by a hypsodont dental crown, the
ready reduced the premolar row and snout, at least as occurrence of the caprine fold, the absence of basal pil-
much as the type population of Capo Figari. The Capo lars in the lower molars, and mandibles with a horizon-
Mannu D4 suid is thus attributed to Sus sondaari. tal ramus that progressively increases in height from the
Sus cf. sondaari has recently been reported in de- diastema to a position under M3 (Plate I, figs. F-M,
posits close to the Plio-Pleistocene transition at Monte Tab.1). The features cited above and the size make these
Tuttavista (Orosei, Eastern Sardinia, Abbazzi et al.
2004). This material is currently under study.
Sus sondaari from Capo Mannu represents the
oldest documentation of the taxon. M2 DAP M2 DT M3 DAP M3 DT

Capo Mannu 13.94 10.37 18.37 12.81

Artiodactyla Owen, 1848 Capo Figari 15.10 14.90 19.40 14.60

15.10 13.80 23.20 14.80
Bovidae Gray, 1821 18.00 14.80 20.60 14.10
17.80 15.00 21.20 14.10
Several fragmentary specimens make the Bovidae Tab. 1 - Measurements (mm) of M2 and M3 of Sus sondaari from
the best represented taxon within the large mammal Capo Mannu Formation and Capo Figari (holotype). Le-
assemblage from the Capo Mannu Fm. Unfortunately gend: DAP=maximum length; DT=maximum width.
Fossil vertebrates from Capo Mannu, with description of a new Testudo species 129

remains comparable to those referred to the Sardinian Bovid remains of small size which do not fully conform

endemic bovid Nesogoral melonii Dehaut (Gliozzi & to the morphology of Nesogoral spp.

Malatesta 1982) which characterises Late Pliocene- Material: Capo Mannu D4 local fauna - CL 137 right hemi-
Early Pleistocene Sardinian endemic faunas (e.g. ``Neso- mandible; Capo Mannu D4 local fauna - CL128, left M3; CL 105, CL
goral complex'' according to Sondaar 2000). 110, two fragments of lower molars.
The referral of the remains from Capo Mannu to
Nesogoral sp. is, however, necessary due to the fragmen- Remarks. The morphology of specimen CL 137
tary state of fossils that hampers detailed comparison. (Plate I, Figs N-P) is different in many respects from
One of the most peculiar features of Nesogoral is that of the remains referred to Nesogoral sp. from Capo
the triangular shape of the hemimandibles, found also in Mannu and from Nesogoral melonii found at the type
the remains from the Capo Mannu formation. This fea- site Capo Figari (Gliozzi & Malatesta 1982). It has a
ture, due to the increased height of M3, occurs as a lower and regularly curved horizontal ramus and, on
parallelism among endemic insular bovids, and is seen the labial side, lacks the protruding angular tuberosity
in Myotragus from the Late Miocene-Holocene of the for the insertion of the masseter muscle, which is pre-
Balearic Islands and in Maremmia from the late Mio- sent in CL127 and GP1. The completely worn teeth do
cene Tusco-Sardinian paleobioprovince. not allow for further comparisons.
The M3 (CL128) and the fragmentary lower mo-
Nesogoral sp.
lars (CL 105, CL 110) are relatively narrower, show less
GP1 M3 19.5 19.1 7.8 5.7 developed stylids, and an almost absent caprine fold
GP1 M2
GP1 M1
- (Fig. 7, Tab. 2), compared to Nesogoral.
CL138 M1 12.8 *13 6.5 6.9 The variability of the above features is poorly
known in the fossil samples of Nesogoral, which neces-
CL127 M2 13.5 12.7 7.1 6.9
CL123a M2 16.4 15.1 14.0 8.1
CL120 M3
Bovidae undet. (small sized)
15.7 14.5 14.1 5.6
sitates separate consideration for the remains described
CL128 M3 21.5 21.1 7.8 27.7 in this section.
Bovidae gen. et sp. indet.
CL129 M3 24.3 24.2 9.6 20.3
NN M2 16.2 * 15 10.4 16.9

Tab. 2 - Measurements (mm) of bovid teeth from Capo Mannu Bovidae gen. et sp. indet. (larger sized)
Formation. Legend: OL= occlusal length; NL= length
at the neck; W= Width; H= height at the metastylid/me- Material: Capo Mannu D4 local fauna - CL129, left M3;
sostyle. * = inferred measurement. CL123b, fragment of upper molar; Not numbered, left M2.

Fig. 7 - Small - sized bovid of uncer-

tain attribution - A, B, C la-
bial, lingual and occlusal
views of CL128 left M3 from
Capo Mannu D4 local fauna.
Bovidae gen. et sp. indet.
(large sized) - D, E, F occlu-
sal, labial and lingual views
of CL129 left M3 from Capo
Mannu D4 local fauna.
130 Abbazzi A., Carboni S., Delfino M., Gallai G., Lecca L. & Rook L.

An M3 and an M2 are clearly distinct in their the contact is covered by Upper Pleistocene MIS 5-2
larger size (Tab. 2); morphologically, they have more sediments, while the sequence of dune units from D2
rounded lobes on the lingual side (Fig. 7) compared to to D7 displays a clear depositional continuity with the
the remains referred to Nesogoral sp. presence of the first-order boundaries. The D1 unit di-
rectly overlying the Mandriola Limestone is made up of
Remarks. The bovid postcranial remains from the a foraminiferal grainstone, whilst D2 is composed of
Capo Mannu D4 local fauna are attributed to two bioclastic and siliciclastic sands. Together with the dif-
groups on the basis of size differences, without any ferent composition of the two units, it can also be ob-
precise systematic attribution: served that D1 shows a higher degree of lithification
compared to D2. Therefore, it is not possible to assign
Smaller sized remains: CL141a, right metacarpal; CL116, me-
tacarpal; CL112, II phalanx; CL104, unciform; CL118, distal fragment a precise age to the end of the accretion of the whole
of humerus; CL135, incomplete tibia; CL122, diaphysis of left tibia; dune complex, or to evaluate the beginning of the ero-
CL139, proximal end of left radius; CL133, fragment of diaphysis of sion phase before the deposition of the Upper Pleisto-
radius; CL112, distal fragment of radius; CL 134, and CL136, frag- cene MIS 5 beach sediments.
ments of pelvis. Nevertheless, in the lack of a clear erosional trun-
Larger sized remains: CL131, incomplete metacarpal; CL 145a
and 145b, fragments of metacarpal; CL146, incomplete humerus; cation, only an insignificant stratigraphic lacuna, unable
CL126, fragment of tuber calcanei; CL126, fragment of diaphysis of to modify substantially the age previously hypothesized
metatarsal; CL 108, proximal fragment of metacarpal. on the basis of the cyclic stratigraphic model, could be
To this undetermined material, can be added a badly crushed present between D1 and D2.
fragment of a bovid skull (CL125). As a result, whilst the transition of the shoreface-
backshore dune (unit D1) of the Mandriola section can
be correlated with the Zanclean-Piacenzian transition,
and the overlying D2-D4 units correlate well with the
The depositional features and the composition of Upper Pliocene, only the thickest of the uppermost
the entire Capo Mannu dune complex suggest particular dune units of the CFM may have reached the Lower
paleogeographic conditions which are very different Pleistocene (Fig. 4).
from the modern exposed and submerged coastal mor-
phology. The dune accumulation may have resulted
from a hot and dry climatic regime with intense aeolian Conclusions

dynamics. The calcareous bioclastic component, as well

as the benthic foraminifers are evidence of a transitional The fossil record of Pliocene vertebrates in Sar-
continental coastal paleoenvironment during sea-level dinia is relatively poorly known. The Capo Mannu D4
high stands. Several reddish sediment layers and dune faunal assemblage allows us to partially highlight a
units very rich in terrigenous siliciclastic components chapter of this Pliocene history. Sondaar (2000) intro-
are probably related to fluctuations in climate towards duced the phrase ``Nesogoral complex'' to refer to an
cooler and wetter regimes, coinciding with regressive assemblage occurring in fissure fillings at Capo Figari
marine eustatic phases. These sediments, likely asso- (Olbia) and Monte Tuttavista (Orosei) dated to the
ciated with a climatic-eustatic cyclicity, may be tenta- latest Pliocene-early Pleistocene. The ``Nesogoral com-
tively analysed according to cyclic stratigraphic models plex'' (cf. also Sondaar & Van der Geer 2002) is char-
to define an upper time limit of the genesis of the entire acterised by the occurrence of endemic species of the
dune complex. Maximum values on the order of 1 Ma murid Raghapodemus (Raghapodemus minor), the bo-
are obtained by correlating the nineteen dune subunits vid Nesogoral, the small suid Sus sondarii, and the
with the astronomical cycles of precession or obliquity endemic macaque Macaca majori. A recent compre-
of the earth's axis (19-23 or 41-53 ky), and the seven hensive study of the extremely large record from the
dune units to the cycle of eccentricity of the earth's Monte Tuttavista faunal complexes allowed for the re-
orbit (97-127 ky), while the minimum values obtained cognition within this ``Nesogoral complex'' of peculiar
are about 0.4-0.5 Ma. On the basis of such a hypothesis, carnivores like the mustelid Pannonictis and the hyae-
the deposition of the Capo Mannu Fm. dune complex, nid Chasmaporthetes melei, as well as a differentiated
starting from the uppermost part of the Mandriola bovid assemblage (Abbazzi et al. 2004; Rook et al.
Limestone, would have been completed before the end 2003; 2004).
of the Pliocene. Although the Capo Mannu D4 mammal assem-
However, field data show that no clear strati- blage does not allow us to derive precise chronological
graphic conformity can be observed between the first constraints, a few considerations can be made. First of
dune unit D1 (containing the Capo Mannu D1 local all, the evolutionary stage of the endemic murid Ragha-
fauna), and the overlying second dune unit D2, because podemus from the Capo Mannu D1 local fauna (for-
Fossil vertebrates from Capo Mannu, with description of a new Testudo species 131

merly known as ``Mandriola''; from an interdune de- representing the oldest known sample of the ``Nesogoral
posit within the basal Capo Mannu Fmt D1 dune unit, complex''.
cfr. Fig. 2), although already endemized (Raghapode- Acknowledgments. For profitable discussion and help with com-
mus azzarolii) shows only incipient modification com- parative collections we are grateful to M. Arca (Nuoro), S. Gentili (Per-
pared to the later species Raghapodemus minor, typical ugia), T. Kotsakis (Rome), M. Palombo (Rome), C. Tuveri (Nuoro), J.
of the ``Nesogoral complex'' as defined by Sondaar van der Made (Madrid). F. de Lapparent de Broin (Paris) kindly offered
her assistance for the identification of the tortoise; D. Righi (Modena)
(2000) and recognized at Monte Tuttavista (Abbazzi discussed the morphology of T. amiatae; E. Razzetti (Pavia) suggested
et al. 2004). relevant literature concerning extant tortoises. We specially thank Dr.
As a matter of fact, given the biochronologic char- Maria Tegas (Lanusei, Sardinia) for the contribution in the discovery of
acterization of Raghapodemus azzarolii (MN 15?; An- the tortoise during the field survey of her degree thesis.
gelone & Kotsakis 2001) and the above discussed more This paper was presented at the 32nd International Geological
likely maximum estimate of 1 Ma for the deposition of Congress, Florence 2004. We thanks M. Chiari (Firenze) for his editor-
ial work on a first version of the manuscript and A. Shabel (Berkeley),
the entire dune complex Ð even in case of a minor M.R. Palombo (Rome) and L. Werdelin (Stockholm) for critical revi-
stratigraphic lacuna between D1 and D2 Ð the age of sion of the manuscript.
the vertebrate remains from the D3-D4 dune units Research supported by Fondi d'Ateneo grants (LR) and Uni-
should be constrained within the Late Pliocene, thus versitaÁ degli Studi di Cagliari (M.I.U.R 60%) grants (SC).


Abbazzi L., Angelone C., Arca M., Barisone G., Bedetti C., Carboni S. & Lecca L. (1985) - Osservazioni sul Pleistocene
Delfino M., Kotsakis T., Marcolini F., Palombo M.R., Medio-Superiore della penisola del Sinis (Sardegna
Pavia M., Piras P., Rook L., Torre D., Tuveri C., Valli occidentale). Boll. Soc. Geol. Ital., 104: 459-477,
A.M.F. & Wilkens B. (2004) - Plio-Pleistocene fossil Roma.
vertebrates of Monte Tuttavista (Orosei, E. Sardinia, Carboni S. & Lecca L. (1995) - Le PlioceÁne de Capo Mannu
Italy), an overview. Riv. It. Paleont. Strat., 110(3): (Sardaigne occidentale): transition marin littoral-con-
603-628, Milano tinental dunaire. C.R. Acad. Sci. Paris, 320, seÂr. II a:
Ambrosetti P., Azzaroli A. & Kotsakis T. (1980) - Mammi- 1203-1210, Paris.
feri del Plio-Pleistocene delle isole. In: I Vertebrati Cherchi A., Marini A., Murru M. & Robba E. (1978) - Stra-
fossili italiani: 243-348, Museo Civico di Storia Na- tigrafia e paleoecologia del Miocene superiore della
turale di Verona, Verona Penisola del Sinis (Sardegna occidentale). Riv. It. Pa-
Amiranashvili N.G. (2000) - Differences in shell morphol- leont. Strat., 84(4): 973-1036, Milano.
ogy of Testudo graeca and Testudo hermanni, based Delfino M. (2002) - Erpetofaune Italiane del Neogene e del
on material from Bulgaria. Amphibia-Reptilia, 21: 67- Quaternario. Doctoral Dissertation Thesis. 382 pp.,
81, Leiden. Modena and Reggio Emilia University.
Amori G., Angelici F.M., Frugis S., Gandolfi G., Groppali Esu D. (1984) - La malacofauna continentale pliocenica di
R., Lanza B., Relini G. & Vicini G. (1993) - Verte- Mandriola (Sardegna occidentale): sistematica e paleo-
brata. In: Minelli A., Ruffo S. & La Posta S. (Eds) - biogeografia. Geologica Romana, 23: 23-50, Roma.
Checklist delle specie della fauna d'Italia. Vol. 110, IV Esu D. & Kotsakis T. (1979) - Restes de VerteÂbreÂs et de
Mollusques continentaux dans le Villafranchien de la
+ 83 pp., Calderini, Bologna. Sardaigne. GeÂobios, 12(1): 101-106, Lyon.
Angelone C. & Kotsakis T. (2001) - Rhagapodemus azzarolii Esu D. & Kotsakis T. (1983) - Les verteÂbreÂs el les mollus-
n. sp. (Muride, Rodentia) from the Pliocene of Man- ques continentaux du Tertiaire de la Sardaigne: paleÂo-
driola (Western Sardinia - Italy). Boll. Soc. Paleont. biogeographie et biostratigraphie. Geologica Romana,
Ital., 40(2): 127-132, Modena. 22, 177-206, Roma.
Batsch G.C. (1788) - Versuch einer Unleitung, zur Kenntnis Francolini L., Lecca L. & Mazzei R. (1990) - La presenza del
und Geschichte der Thiere un Mineralien. Jena, Ack- Pliocene inferiore nella piattaforma continentale della
ad. Buchh., 1: i-vi, 1-528, Iena. Sardegna occidentale. Atti Soc. Tosc. Sci. Nat., Mem.,
Beccaluva L., Civetta L., Macciotta G. & Ricci C.A. (1985) - Ser. A, 97: 93-111, Pisa.
Geochronology in Sardinia: results and problems. Gliozzi E. & Malatesta A. (1982) - The Quaternary Goat of
Rend. Soc. Ital. Mineral. Petrogr.: 40, 57-72, Varese. Capo Figari (Northeastern Sardinia). Geologica Ro-
Brongniart A. (1800) - Essay d'une classification naturelle mana, 19 (1980); 295-347, Roma.
des Reptiles. Bull. Soc. philomat., 3, 2, 11 (=35): 81- Gray J.E. (1821) - On the natural arrangement of vertebrose
82; 12 (=36): 88-91, Paris. animals. London Medical Repository, 15; 296-310,
Brookfield M.E. (1977) - The origin of bounding surfaces in London.
ancient aeolian sandstones. Sedimentology, 24: 303- Guyot Jackson G. (ed.) (2004) - NumeÂro speÂcial - Testudo.
322, Oxford. Manouria, 7(22): 1-52, Mezzavia.
132 Abbazzi A., Carboni S., Delfino M., Gallai G., Lecca L. & Rook L.

Hervet S. (2001) - EÂtude du peuplement chelonian de la Rend. Sem. Fac. Sc. Univ. Cagliari, suppl. vol. 43:
Corse aÁ partir de la repartition des tortues fossils de 305-320, Bologna.
la MeÂditerraneÂe occidentale. Bull. Soc. Sci. Hist. Nat. Rook L., Abbazzi L., Angelone C., Arca M., Barisone G.,
Corse, 696-697: 147-163, Bastia. Bedetti C., Delfino M., Kotsakis T., Marcolini F., Pa-
Hervet S. & Salotti M. (2000) - Pleistocene turtles from lombo M.R., Pavia M., Piras P., Torre D., Tuveri C.,
Castiglione (Oletta, Corsica) and evidence of indigen- Valli A. & Wilkens B. (2003) - Osservazioni prelimi-
ous population in Corsica. C.R. Acad. Sci. Paris, ser. nari sui vertebrati fossili Plio-Pleistocenici del Monte
II a, 330(9): 645-651, Paris. Tuttavista (Orosei, Sardegna). Sardinia Corsica et Ba-
Kotsakis T. (1980) - Osservazioni sui Vertebrati quaternari leares Antiqvae, 1: 11-29, Pisa-Roma.
della Sardegna. Boll. Soc. Geol. Ital., 99: 151-165, Rook L., Ferretti M.P., Arca M. & Tuveri C. (2004) - Chas-
Roma. maporthetes melei n. sp., an endemic Hyaenid (Car-
Lanza B. & Corti C. (1993) - Erpetofauna italiana: ``acqui- nivora, Mammalia) from the Monte Tuttavista fissure
sizioni'' ed estinzioni nel corso del novecento. Supp. fillings (Late Pliocene to Early Pleistocene; Sardinia,
Ric. Biol. Selvaggina, 21: 5-49, Bologna. Italy). Riv. It. Paleont. Strat., 110(3); 639-646, Milano.
Lapparent de Broin de F. (2000) - African chelonians from Sondaar P.Y. (2000) - Early Human Exploration and Exploi-
the Jurassic to the present: phases of development and tation on Islands. Tropics, 10: 203-230, Kagoshima.
preliminary catalogue of the fossil record. Palaeont. Sondaar P.Y. & Van der Geer A. (2002) - Plio-Pleistocene
afr., 36: 43-82, Johannesburg. vertebrate faunal evolution on Mediterranean islands,
Lapparent de Broin de F. (2001) - The European turtle fauna compared to that of the Paleartic mainland. Ann.
from the Triassic to the Present. Dumerilia, 4: 155- GeÂol. Pays HelleÂniques, 39(A), 165-180, Athens.
217, Paris. Tegas M. (2000) - Contributo alla stratigrafia del Pliocene,
Lapparent de Broin de F. (2002) - A giant tortoise from the Pleistocene ed Olocene del Sinis nord-occidentale.
Late Pliocene of Lesvos Island (Greece) and its pos- Unpublished Thesis, UniversitaÁ degli Studi di Ca-
sibile relationships. Ann. GeÂol. Pays HelleÂniques, gliari, 144 pp.
39(A): 99-130, Athens. Van der Kuyl A.C., Ballasina D.L.P., Dekker J.T., Maas J.,
Lecca L. & Carboni S. (2007) - The Tyrrhenian section of Willemsen R.E. & Goudsmit J. (2002) - Phylogenetic
San Giovanni di Sinis (Sardinia): stratigraphic record relationships among the species of the genus Testudo
of an irregular single ligh stand. Riv. It. Paleont. (Testudines: Testudinidae) inferred from mitochon-
Strat., 113(3): 509-523, Milano. drial 12S rRNA gene sequences. Mol. Phylog. Evol.,
Lecca L., Carboni S., Scarteddu R., Pisano S., Sechi F. & 22: 174-183, Amsterdam.
Tilocca G. (1986) - Schema stratigrafico della piatta- Van der Made J. (1988) - Sus nanus nov. sp., a Pliocene dwarf
forma continentale occidentale e meridionale della pig from Capo Figari (Northeastern Sardinia). Boll.
Sardegna. Mem. Soc. Geol. It., 36: 31-40, Roma. Soc. Paleont. Ital., 27(3): 367-378, Modena.
Linnaeus C. (1758) - Systema Naturae. Ed. 10, Vol. I, Re- Van der Made J. (1999) - Biogeography and stratigraphy of
gnum Animale. Holmiae (Stockholm), L. Salvii, 1- the Mio-Pleistocene mammals of Sardinia and the de-
824, Stockholm. scription of some fossils. In: Reumer J.W.F & De Vos
Owen R. (1848) - Contributions to the History of British J. (Eds) - ``Elephants have a snorkel! Papers in honour
Fossil Mammals. 71 pp., Taylor, London. of Paul Y. Sondaar'', Deinsea, 7: 337-360, Rotterdam.
Palombo M.R., Valli A.M.F., Arca M. & Tuveri C. (2006) - A Van der Made J. (2005) - The fossil endemic goat Nesogoral
new bovid, Asoletragus gentryi n. gen. et sp., from cenisae n. sp. from Campidano, Sardinia - Cursorial
Monte Tuttavista (Orosei, Eastern Sardinia, Italy). adaptations in insular environment. In: Alcover J.A.
Riv. It. Paleont. Strat., 112(3): 459-471, Milano & Bover P. (Eds) - ``Insular Vertebrate Evolution: the
Pecorini G., Rage J.-C. & Thaler L. (1974) - La Formation Palaeontological Approach'', Monografies de la Soci-
continentale de Capo Mannu, sa faune de VerteÂbreÂs etat d'Historia Natural de les Balears, 12: 347-368,
plioceÁnes et la question du Messinien en Sardaigne. Palma de Mallorca.

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